Global Patterns of Guild Composition and Functional

Diversity of Spiders
Pedro Cardoso1,2*, Stano Pekár3, Rudy Jocqué4, Jonathan A. Coddington1
1 Smithsonian Institution, National Museum of Natural History, Washington DC, United States of America, 2 Azorean Biodiversity Group - CITA-A, Universidade dos Açores,
Angra do Heroı́smo, Portugal, 3 Department of Botany and Zoology, Faculty of Science, Masaryk University, Brno, Czech Republic, 4 Musée Royal de l’Afrique Centrale,
Tervuren, Belgium

Abstract
The objectives of this work are: (1) to define spider guilds for all extant families worldwide; (2) test if guilds defined at family
level are good surrogates of species guilds; (3) compare the taxonomic and guild composition of spider assemblages from
different parts of the world; (4) compare the taxonomic and functional diversity of spider assemblages and; (5) relate
functional diversity with habitat structure. Data on foraging strategy, prey range, vertical stratification and circadian activity
was collected for 108 families. Spider guilds were defined by hierarchical clustering. We searched for inconsistencies
between family guild placement and the known guild of each species. Richness and abundance per guild before and after
correcting guild placement were compared, as were the proportions of each guild and family between all possible pairs of
sites. Functional diversity per site was calculated based on hierarchical clustering. Eight guilds were discriminated: (1)
sensing, (2) sheet, (3) space, and (4) orb web weavers; (5) specialists; (6) ambush, (7) ground, and (8) other hunters. Sixteen
percent of the species richness corresponding to 11% of all captured individuals was incorrectly attributed to a guild by
family surrogacy; however, the correlation of uncorrected vs. corrected guilds was invariably high. The correlation of guild
richness or abundances was generally higher than the correlation of family richness or abundances. Functional diversity was
not always higher in the tropics than in temperate regions. Families may potentially serve as ecological surrogates for
species. Different families may present similar roles in the ecosystems, with replacement of some taxa by other within the
same guild. Spiders in tropical regions seem to have higher redundancy of functional roles and/or finer resource partitioning
than in temperate regions. Although species and family diversity were higher in the tropics, functional diversity seems to be
also influenced by altitude and habitat structure.

Citation: Cardoso P, Pekár S, Jocqué R, Coddington JA (2011) Global Patterns of Guild Composition and Functional Diversity of Spiders. PLoS ONE 6(6): e21710.
doi:10.1371/journal.pone.0021710
Editor: Michael Somers, University of Pretoria, South Africa
Received March 2, 2011; Accepted June 6, 2011; Published June 29, 2011
This is an open-access article, free of all copyright, and may be freely reproduced, distributed, transmitted, modified, built upon, or otherwise used by anyone for
any lawful purpose. The work is made available under the Creative Commons CC0 public domain dedication.
Funding: PC was supported by Fundação para a Ciência e Tecnologia (SFRH/BPD/40688/2007). SP was supported by project no. 0021622416 of the Ministry of
Education, Youth and Sports of the Czech Republic. JAC was supported by various grants from the Neotropical Lowlands, Biodiversity of the Guianas, and
Scholarly Studies Programs of the Smithsonian Institution, the Carlsberg Foundation, and NSF grants EAR-0228699, DEB-9707744, and DEB-9712353. The funders
had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
Competing Interests: This work was partly supported by The Carlsberg Foundation. This does not alter the authors’ adherence to all the PLoS ONE policies on
sharing data and materials.
* E-mail: pcardoso@ennor.org

Introduction environment, independently of the specific taxonomic composi-

Hutchinson [1] was the first to suggest that species were limited to
ecological niche boundaries by competing species [2]. Groups of
competitors, or ‘‘a group of species that exploit the same class of
environmental resources in a similar way’’, were later called guilds by
Root [3,4]. In the meantime, many different definitions of guilds were
used, in a relatively loose way [5,6]. The currently most accepted
definition characterizes ecological guilds as non-phylogenetic groups
of species that share one or a series of important resources [7].
Parallel to guilds, functional groups were defined as groups of
species that have the same function in the ecosystem, providing the
same ecosystem services. Although guilds and functional groups
are different concepts, with the first focusing on resource sharing
and the latter focusing on ecosystem processes, the groups formed
by both approaches often overlap [4,7,8]. Guild members may
have similar functional roles in the communities, in which case
both terms define the same set [7].
The definition and study of guilds is especially useful if they
respond in roughly the same way to similar changes in the
tion. Studying ecological guilds (or functional groups) can be useful
to investigate assemblage response to climate change [8–10],
habitat disturbance [11,12], management [13] among many other
areas [6].
The study of guild structure implies its quantification.
Functional diversity is one of the most important parameters used
to explain how ecosystems work and adapt to change [14,15]. In
order to quantify guild and functional group diversity, a number of
complex and precise measures have been developed during the
latter decade. Using total dendrogram branch length to measure
functional diversity was first proposed by Petchey, Gaston [16] as a
more useful measure than simply counting the number of guilds or
functional groups [14].
Higher trophic levels have been repeatedly found to be
especially sensitive to environmental change, either because they
operate at a larger spatial scale than other groups, becoming more
sensitive to, e.g., fragmentation, or because they are subject to the
same factors as lower trophic levels as well as being strongly
dependent on lower trophic groups and their changes, thus

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experiencing a synergistic effect [8]. Spiders therefore ought to be
a good indicator taxon to reflect ecological change. They are in
fact the main arthropod predators in many biomes and habitat
types. Additionally, they have already been suggested to be an
ideal group for predicting extinction debt in other taxa due to
habitat destruction [17]. Classifying spiders into guilds seems
therefore useful to future studies of ecological change in all kinds of
biomes and habitats.
Several authors have tried to define spider guilds by using
foraging strategies to predict arthropod prey group as the shared
resource [18,19]. Flying arthropods are mainly captured by
different types of webs, epigean arthropods by wandering spiders
or tube web hunters, arboreal arthropods by sheet webs, etc.
Therefore, although many guild classification systems exist for
spiders, these are usually based solely on foraging strategy,
although different strategies may be directed towards similar prey
and similar strategies may be directed towards different prey.
Based on comprehensive spider inventories of a number of
forest sites in different parts of the world, our first objective was to
define spider guilds that can be applied to all extant families
worldwide. Comparing spider assemblages in different regions is
possible if guild and functional diversity patterns are global, even if
taxonomic composition is disparate [19]. Because spiders are
among the most abundant and diverse predators in all kinds of
terrestrial biomes worldwide and because predators are predicted
to be especially sensitive to ecological change, it is important to
define guilds applicable at a worldwide level and to verify if guilds
and functional diversity are potentially useful to make comparisons
among taxonomically disparate assemblages. We certainly realize
that the current scope of spider families as presently defined in
some cases is so broad that one family may include various guilds,
functional groups and foraging strategies, but it nevertheless seems
worthwhile to investigate the possibility and to assess critically its
success. This is the first time such a goal is attempted at a global
scale for spiders and, to our knowledge, for any invertebrate
group.
The second objective was to test the hypothesis that guilds
defined at family level are good ecological surrogates of guilds
defined at the species level. Guild classification should ideally be
made at the species level, because each species usually has a
uniform behavior, which may be different from any other, even
closely related, species [19]. However, it is impossible to assign
guilds to all spider species, or even genera. Currently 110 families
(two new families have recently been recognized), 3,821 genera,
and more than 42,000 spider species are known [20]. Rates of
description are high, limited mainly by the taxonomists available,
and based overwhelmingly on museum specimens without
ecological data. The behavior of probably 90% of the described
species is unknown and, if necessary, inferred from the genus or
family to which they belong. Phylogeny is probably the best
predictor of ecology, but even at the family level spider phylogeny
is not robustly known. Intrafamilial phylogenies are scarce,
equivocal, and almost universally based on dramatically incom-
plete samples. About 75% of the genera contain five or fewer
species and many are known only from the original literature
description. Roughly half of the species descriptions are pre-1940
and contain only morphological information. Nevertheless,
experienced araneologists expect to identify animals in the field
to family from behavior, habits, habitat, and appearance, so
‘‘ecology’’ is relatively predictable at coarse scale. Therefore, using
higher taxa surrogates at the family level may be as justified for
guild classification and functional diversity quantification (ecolog-
ical surrogacy) as it is for taxonomic diversity quantification
(taxonomic surrogacy) [21]. Given that family members tend to
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Guilds and Functional Diversity of Spiders
have similar lifestyles [18,22,23], we hypothesize that using family
surrogates can be an appropriate strategy.
The third objective was to compare the taxonomic (family) and
guild composition of spider assemblages from different parts of the
world. Taxonomic composition of assemblages varies wildly
between sites in different biomes. Usually no native species and
only a few genera are shared between temperate and tropical
forests. Families may be exclusive to particular regions; richness or
abundance of families usually differ. However, spiders and their
role as one of the main predator taxa in all terrestrial ecosystems
are ubiquitous. If ecosystem services are similar worldwide, with all
ecosystems needing the same functional components indepen-
dently of what taxa perform which tasks, different communities
may present similar guilds in similar proportions even if the
taxonomic composition differs. This similarity of guilds in different
regions forms the basis for the apparent convergence of distant
assemblages [5,24]. We therefore hypothesize that guild compo-
sition is more stable, i.e. constant in proportions, at global scales
than taxonomic composition.
Our fourth objective was to compare the taxonomic and
functional diversity of spider assemblages from different parts of
the world. Tropical regions are known to be major hotspots of
biodiversity, with species richness reaching its peak for most
animal taxa. A number of alternative or complementary
hypotheses have been suggested to explain this almost universal
pattern, from tropical climates being older and historically larger,
allowing more opportunity for diversification [25] to geometric
constraints on species richness [26,27]. However, higher richness
may not equate to higher functional diversity, because species may
have partly redundant roles and/or establish a finer resource
partitioning. Given that co-occurring taxonomically similar species
tend to diverge in their functional roles in order to avoid
competition, poorer assemblages may present species that occupy
the available niches as thoroughly as the species in richer
assemblages. We hypothesize that although taxonomic (family)
richness is considerably higher in the tropics, differences in
functional diversity will be much less, at least between different
biomes, and will show higher redundancy and/or finer resource
partitioning in lower latitudes.
Our fifth and final objective was to relate functional diversity
with habitat structure and complexity. If functional diversity
remains mostly similar across latitudes and biomes, other
explanations must be sought to understand differences between
sites. Habitat structure may be responsible for such differences. A
site with major vegetation complexity can present more variety of
prey or simply more opportunities for spiders to build snares and
retreats. We hypothesize that functional diversity is positively
related with habitat complexity, with more complex habitats being
more functionally diverse.
Materials and Methods
Study sites and sampling procedures
This study is based on datasets from work on spider diversity
over the last twenty years under diverse conditions and objectives.
Seven forest sites were chosen in different regions of the world to
provide a reasonable synopsis of global spider diversity (Table 1).
Sampling followed the semi-quantitative design of Coddington
et al. [28], with different effort per method at each site (see
references in Table 1). In this kind of sampling, each sample
represented one method applied for 1 h of active, continuous
collecting (i.e. including time required to transfer the specimens to
a vial, but excluding interruptions). Semi-quantitative sampling as
applied for this work, especially if optimized, was recently found to
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Table 1. Overview of studied sites and respective biogeographical, ecological and spider assemblage characteristics.

Estimated
species
Altitude Family Species Number of richness
Site Climatic region Habitat type (m) richness richness individuals (Chao 1) Reference

USA Temperate Mesic hardwood forest 800 17 76 1228 108 [29]

Portugal Temperate/ Mixed English oak (Quercus 650 25 117 1795 142 [30]
(Gerês) Mediterranean robur L.) and Pyrenean oak
(Quercus pyrenaica Willd.)
woodland
Portugal Mediterranean Cork oak (Quercus suber L.) 60 26 93 1473 106 [31]
(Arrábida) woodland
Guyana Tropical Lowland moist forest 240 31 228 2934 234 [32]
Cameroon Tropical Mid-elevation moist forest 800 32 218 1407 310 J.A. Coddington et al.,
unpublished data
Tanzania Tropical High-elevation moist forest 1850 31 120 2330 155 [33]
Madagascar Tropical Mid-elevation moist forest 1000 33 291 3167 334 J.A. Coddington et al.,
unpublished data

doi:10.1371/journal.pone.0021710.t001

be extremely efficient and capable of guaranteeing maximum
richness with minimum effort but still allowing comparability of
sites by using a standard set of methods and effort per method and
time of day [34,35]. Sampling was made by different teams at
each site, however, all teams had a mix of experienced and
inexperienced collectors and collecting experience with semi-
quantitative sampling was previously found to be relatively less
important than method and time of day for sampling efficiency
[30,31,36]. We limited the sites to forests, from temperate to
tropical, to reduce variance. Savannah or similar open habitats
require a different set of sampling methods [34,35,37]. In
Cameroon we had to lump data from two different plots in order
to increase sample number, however, 84% of the data came from
a single plot. The higher elevation locales in the tropics had lower
canopies and in general a simpler habitat structure than low
elevation sites.
Because for every site we had a different number of samples per
method, to guarantee the comparability of datasets for each one,
we considered 32 samples of each of three methods:
Aerial searching - Hand collection with pooter, vial, forceps or
brush from knee level to as high as the collector could reach.
Beating - Branches of trees and other vegetation were beated
with a wooden stick while holding a 1-m square beating tray
underneath to catch the falling specimens.
Ground searching - Hand collection from ground level to knee
height.
All three methods were applied equally during day and night
(spiders are mainly nocturnal), i.e., 16 hours of sampling per
method/time of day in each site. In the few cases when enough
samples of one method/time of day combination were not
available (diurnal aerial sampling in Guyana (12 samples) and
nocturnal beating in Cameroon (9 samples)) the missing data were
substituted by samples of the same method but a different time.
Ecological data
The definition of guilds should be based on ecological
characteristics of species (or higher taxa) that determine resource
sharing. As mostly generalist predators of arthropods, the most
important resource for spiders is arthropod prey, and their most
important distinctive characteristics probably are their foraging
method, the range of prey they hunt, vertical stratification,
circadian activity, body size and phenology. Body size and
phenology within spider families are extremely variable world-
wide, and present wide disparities in these traits. Given the broad
scale and exploratory character of this study, we therefore have
not considered such traits, although they are no doubt important
in structuring assemblages at a local scale. In this work we used
information on foraging strategy (type of web and method of active
hunting), prey range (either stenophagous or euryphagous),
vertical stratification (ground or vegetation) and circadian activity
(diurnal or nocturnal) (Table S1).
Data for each family was collected from a number of sources.
We used the general characteristics of families [18,22,23],
acknowledging that exceptions in many cases are inevitable at
such a large taxonomic and geographic scale. In a few families in
which relatively large numbers of species clearly have evolved
distinct lifestyles (i.e. not just the exception), we separated the
families into sub-families and classified each accordingly. These
sub-families are hereafter treated as families. When families were
largely unknown, we used the characteristics of a particular species
in the family for which the behavior was known. Additionally, we
analyzed data from a number of exhaustive samples combining
methods targeted towards different vertical strata and times of day
[28–33,36,37]. Such data allowed inferring on the vertical
stratification and circadian activity of families. All characteristics
were evaluated only in a binary way (Table S1).
Definition of guilds
The definition of guilds can be made a priori, based on certain
characteristics thought to be especially important [8,38–40], or a
posteriori, using quantitative methods to find natural groups [41].
The latter reduces the subjectivity of guild placement. However,
resulting groups may seem less natural to the experienced
researcher [42]. Among quantitative methods for the definition
of guilds, we may include nearest-neighbor variance in overlap
[43], multivariate analysis [44], clustering algorithms [18,45,46],
psychometric analysis [47] and bootstrap randomization algo-
rithms [41]. Probably the most commonly used are hierarchical
clustering algorithms. For these spider data at the global level we
opted to use the UPGMA with Sørensen similarity measure
analysis, as these methods were already used for previous spider
classifications [18].

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Any dichotomous clustering method, such as UPGMA, divides
taxa in 2 to n groups, with n equal to the number of taxa. The
definition of guilds implies this division. However, it may be
difficult to determine where the cut-off of each branch in the tree
representing a separate guild should be made. We decided a priori
to use a set of rules to achieve important practical goals. Firstly,
enough guilds should be recognized to allow useful comparisons of
sites based on guild proportions. Secondly, guilds should not be so
numerous that the smallest would contain only one or very few
families. Thirdly, the guilds should be as homogenous as possible,
although some exceptions were inevitable, for example due to
major divergent lifestyles inside a single family. Although these
rules require some subjectivity, their application after quantitative
analysis mitigated this weakness.
Finally, in order to verify that the defined guilds were
statistically supported, we performed an analysis of similarity
(ANOSIM). This statistic employs a randomization technique that
compares the within and between group similarity of elements as
measured by the Bray-Curtis index. Because we used presence/
absence data, this index was equivalent to the Sørensen index, also
used for building the tree, making both statistical approaches fully
comparable.
Sampling effort
Richness comparisons of assemblages must always be made
cautiously; in particular sampling completeness should not differ. If
some guilds were differentially sampled by the methods employed,
different completeness values would compromise direct compari-
sons. We first calculated for each site each guild’s estimated richness
with the Chao1 estimator [48] and calculated completeness as the
observed to estimated richness ratio. However, the Chao estimates
were far from reliable, and the completeness variance of the
different guilds belonging to each site was very large (results not
shown). The completeness values were therefore unreliable. As an
alternative to completeness, we estimated the final slopes of guild
species richness accumulation curves for all guilds at each site. All
curves were sample-based, randomized 1000 times and rescaled to
individuals as suggested by Gotelli and Colwell [49]. The final
slopes of curves were calculated as:
Slope~(Sa - Sa-1 )=(na - na-1 )
where Sa = total number of species; Sa-1 = number of species after
adding the next to last sample; na = total number of individuals;
na-1 = number of individuals after adding the next to last sample.
The slopes at the end of the accumulation curves for all guilds did
not differ significantly (Kruskal-Wallis test: H7,64 = 10.978;
p = 0.140). Thus all guilds were sampled at a similar rate during
the accumulation process such that higher sampling completeness at
some sites did not influence the proportion of species richness per
guild at each site.
Higher taxa surrogacy
Our datasets included mainly tropical assemblages, for which a
large part of the morphospecies could not be assigned to known
species or even genera. Therefore, to test the higher taxa surrogacy
hypothesis, we used only the two Portugal datasets (Table 1) for
which substantial information about most species was available.
For both datasets, which had intermediate family and species
richness values, we calculated the number of species and
individuals per guild as identified by the methods above. We then
corrected guild placement using the knowledge we had on each
species, and again calculated the number of species and individuals
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Guilds and Functional Diversity of Spiders
per guild. To test the surrogacy, we used a bootstrapping
procedure to evaluate if the richness and abundance differed
statistically for each guild. We also tested the correlation of
richness and abundance per guild before and after correction with
the Spearman rank statistic.
Taxonomic and guild composition
To compare the taxonomic and guild composition between sites
we compared the proportions of each guild and family between
every possible pair of sites using the Spearman rank correlation
index. If guilds have relatively similar proportions worldwide, their
correlations should be higher than the respective family proportion
correlations. For these analyses we used only the eight most rich or
abundant families on average for all sites, so that rare families did
not artificially decrease rank correlation values of the family
comparisons.
Functional diversity
Although many options exist for calculating functional diversity
(FD) [15,50], none are optimal in all cases [51]. For consistency we
used the same UPGMA tree used to define guilds. FD was
calculated as the sum of lengths of the branches connecting all
families observed in a particular site [16]. The complete tree with
all 108 spider families was used for all sites.
Functional diversity depends strongly on taxa richness [16],
because more taxa imply more branches in the tree. Richness, in
turn, strongly depends on the number of observed individuals. We
therefore resampled the data for each site by randomly selecting
1000 individuals and calculating the FD value of the resulting tree.
This resampling was made 1000 times per site, allowing obtaining
95% confidence limits calculated as the respective 0.025 and 0.975
percentiles. All calculations were made with Java software written
specifically for this work (available from the first author by
request).
Results
Definition of guilds
As mentioned above, we recognized subgroups within four
families (Amphinectidae, Desidae, Dictynidae and Linyphiidae)
because their subfamilies exhibited disparate strategies, and
treated these as equal to families. A total of eight guilds could be
discriminated from the UPGMA analysis (Fig. 1, Table S1): (1)
sensing web weavers, (2) sheet web weavers, (3) space web weavers,
(4) orb web weavers, (5) specialists, (6) ambush hunters, (7) ground
hunters and (8) other hunters. The ANOSIM analyses supported
the recognized guilds (global R = 0.917, p,0.001; 0.695,R,1,
p,0.001 in all cases). Guilds ranged in size from relatively large
(ground hunters, 26 families) to small (ambush hunters, six
families). As these separated from other families relatively deeply
in the tree, their discrimination seemed justified.
Higher taxa surrogacy
Out of 173 species and 3268 specimens in the Portuguese
datasets, 27 species (16%) and 358 specimens (11%) were
incorrectly attributed to a guild by family surrogacy. These
included stenophagous species that belong to families where
stenophagy is not very common (e.g., the theridiid ant specialists
Dipoena and Euryopis, the araneophagic jumping spider Cyrba, the
gnaphosid ant specialists Callilepis and Nomisia) or generalist species
in specialist families (e.g., Harpactea and Rhode in Dysderidae). Also,
some higher stratum species occur in typically ground hunting
families (e.g., Echemus and Scotophaeus in Gnaphosidae), and hunting
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Figure 1. Hierarchical clustering of families. Results for the UPGMA analysis with Sørensen index of dissimilarity applied to the ecological
characteristics of spider families. Names of eight distinguished guilds are shown.
doi:10.1371/journal.pone.0021710.g001

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species occur in some web-building families (e.g. Pisaura in
Pisauridae), etc.
Bootstrapping confirmed these differences in numbers of species
and individuals per guild, with space web weavers presenting
lower values and specialists higher values after correcting species
guild placement (Fig. 2). Statistically significant differences were
also found for ambush hunters’ richness and ground hunters’
abundance in both sites, as well as ground hunters’ richness in
Gerês and ambush hunters’ abundance in Arrábida.
However, even given significant differences in richness and
abundance of some guilds before and after correction, the
Spearman rank correlation of uncorrected vs. corrected guilds
was invariably high (0.833,R,0.970; n = 8; 0.00007,p,0.01 in
all cases).
Taxonomic and guild composition
In four cases, the space weavers were the richest guild (between
24 and 35% of species; Fig. 3). In Guyana orb weavers were the
richest, at 35% of species. In USA and Portugal (Arrábida), the
‘‘other hunters’’ guild was the richest with 39% and 28% of species
respectively. Theridiids were richest in six sites, with 12 to 29% of
species (Fig. 2). The exception occurred in the USA where
Linyphiidae (21%) and Araneidae (20%) dominated.
In all cases except the USA and Guyana the most abundant
guilds were the space web weavers, with 33 to 55% of the
individuals captured (Fig. 3). In both exceptions the orb weavers
were most abundant (39 and 40% respectively). Theridiidae was
most abundant in five sites, with 26 to 41% of the individuals
(Fig. 3). However, in the USA Theridiidae was only 9% of the total
abundance, with Araneidae (34%), Linyphiidae (21%) and
Agelenidae (17%) being more abundant. In Tanzania, the
Pholcidae were most abundant (44%), followed by Linyphiidae
(15%), and Theridiidae (10%).
In 19 out of 21 pairwise comparisons of sites (90%) the
correlation of guild species richness was higher than the
correlation of family species richness (Table 2). Significant
correlations were found for guilds in 19 cases (90%), but only in
11 cases (52%) for families. In 20 out of 21 pairwise comparisons of
sites (95%) the correlation of guild abundance was higher than the
Figure 2. Guild richness and abundance per site with family and species guild classifications. Percentage of species and individuals in
each of two sites in Portugal belonging to each guild according to the respective family and species classifications (family classification reflects the
predominant guild in the family (Fig. 1), while species classification represents the true guild of the species).
doi:10.1371/journal.pone.0021710.g002
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Guilds and Functional Diversity of Spiders
correlation of family abundance (Table 2). Correlations were
significant in 19 cases (90%) for guilds, but only in three cases
(14%) for families.
Functional diversity
Family richness was higher in the tropics than in temperate
regions, and the Mediterranean sites in Portugal were intermediate
(Fig. 4). Functional diversity presented a different pattern. The
USA site and the northernmost site in Portugal (Gerês) had the
lowest FD, but the more southern Portuguese site (Arrábida) and
some tropical sites had similar values (Fig. 4). Families in less rich
sites apparently filled the functional tree almost as completely as
families in richer sites, similarly absorbing the available resource
space.
Discussion
Separating species into guilds can be as simple as grouping taxa
according to trophic level [6]; or as complex as studying all the
relationships between taxa in a multidimensional matrix [43]. The
approach taken depends on: (1) the objectives of the study (the
level of detail depends on the information needed to answer
particular questions); (2) the spatial scale of the study (a worldwide
guild classification aims at more general patterns than those at the
assemblage level); (3) the taxonomic scale (classifying species
requires different data than genera or families, whose species may
perform different roles in the same assemblage); (4) the data
reasonably available (quantitative data about all taxa and
interactions is feasible only for single communities). As we
attempted to characterize a megadiverse group at a global scale,
our approach was designed to test questions at this level. In any
case, most tropical spider species are undescribed and discrimi-
nated only as morphospecies. Moreover, ecological characteristics
(foraging strategy, prey range, vertical stratification and circadian
activity) for those species which are identified at specific or generic
levels, are usually unknown. In such cases, family or even genus
characteristics are available to aid in guild assignment. At a global
level for spiders, families, and occasionally groups within families,
are the most practical basis for guild classification.
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Figure 3. Percentage of species and individuals per family and guild in each studied site.
doi:10.1371/journal.pone.0021710.g003

Table 2. Pairwise Spearman rank correlations of family and guild richness and abundance of studied sites.

Portugal Portugal
USA (Gerês) (Arrábida) Guyana Cameroon Tanzania Madagascar

Richness
USA 1 0.814* 0.874** 0.777* 0.635 0.802* 0.599
Portugal (Gerês) 0.122 1 0.857** 0.778* 0.833* 0.881** 0.857**
Portugal (Arrábida) 0.299 0.724* 1 0.850** 0.881** 0.976*** 0.810*
Guyana 0.216 0.834** 0.801* 1 0.898** 0.826* 0.874**
Cameroon 0.180 0.700 0.633 0.946*** 1 0.905** 0.976***
Tanzania 0.619 0.732* 0.778* 0.814* 0.755* 1 0.833*
Madagascar 20.238 0.708* 0.611 0.850** 0.886** 0.524 1
Abundance
USA 1 0.762* 0.690 0.762* 0.762* 0.762* 0.643
Portugal (Gerês) 0.725* 1 0.952*** 0.905** 1.000*** 0.952*** 0.952***
Portugal (Arrábida) 0.691 0.852** 1 0.786* 0.952*** 0.929*** 0.929***
Guyana 0.123 0.515 0.158 1 0.905** 0.762* 0.905**
Cameroon 20.122 0.439 0.193 0.802* 1 0.952*** 0.952***
Tanzania 0.220 0.146 0.133 0.479 0.500 1 0.833*
Madagascar 20.146 0.390 0.205 0.479 0.548 20.143 1

Family below and guild above diagonals. For family richness and abundance we only used the eight most rich or abundant families on average for all sites.
*p,0.05;
**p,0.01;
***p,0.001.
doi:10.1371/journal.pone.0021710.t002

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Figure 4. Average family richness and functional diversity per site (after resampling for 1000 individuals). Letters represent statistically
different (at a,0.05) groups as determined by the resampling procedure.
doi:10.1371/journal.pone.0021710.g004
As expected, the most important characteristic for defining guild
placement was foraging strategy. Web type or hunting methods
primarily determine the division of spider families into guilds, as
was previously recognized by different authors dealing with this
taxon [18,19]. Hunting spiders could be further divided according
to the vertical stratum preferred; stenophagous families constituted
a separate guild. The eight guilds now proposed are partly
coincident with previous classifications dealing with fewer families
in smaller areas. Uetz and colleagues [18], studying crops in the
USA, also proposed eight guilds. These authors, however,
included Linyphiids in their own wandering sheet/tangle weavers
guild and many families that we denominate as ‘‘other hunters’’
were considered as either foliage runners or stalkers. Dias and
colleagues [19], studying Neotropical spiders, further refined many
of the guilds in diurnal and nocturnal. In our global study,
circadian activity was not decisive for guild placement. Circadian
activity, along with phenology and body size, could however be
used in smaller-scale or species-based studies. Species hunting in
different times of day or seasons or having different body sizes
probably are not sharing resources.
No previous study considered the problem of stenophagy and
specialization of prey. It may be important to recognize in guild
placement that specialist taxa have little overlap in resource sharing
with other species. In that sense, the specialists’ guild is not even a
true guild, but a cluster of species that, by specializing in one or very
few prey, are not directly competing with any large group of species.
Some families may seem to be at odds with the common
perception of where they belong in the tree. This may result from
lack of knowledge of the biology of tropical as compared to
temperate species, or that different species have indeed developed
different hunting strategies. For example, scytodids are usually
regarded as ground hunters in the Holarctic, but they predom-
inantly hunt on vegetation in the tropics. Deinopids are specialized
orb weavers whose web is modified to ambush prey. We therefore
characterize deinopids as ambush predators rather than as typical
orb weavers.
In this work we assumed that guilds are largely conserved within
families and that this conservatism was valid at a global level. The
first assumption was tested with the Portugal datasets; our results
supported the use of families as a surrogate for species guild
classification. We could not test the second assumption because
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Guilds and Functional Diversity of Spiders
many individuals are not identified to species or genera in most
tropical datasets. The biology of named tropical species moreover
is usually unknown. However, current knowledge generally
indicates that confamilial spider species tend to have the same
lifestyle [23]. This tendency supports the application of these
findings and guild classification to global biomes and habitats,
although the validity of family surrogacy should be tested
whenever possible to confirm the results here obtained. Even
where direct knowledge of species permits a refined classification
of guilds in a particular study, using a standard set of guilds and
guild denominations will facilitate comparison of studies per-
formed by different teams with different objectives as long as the
methodology used to obtain data is comparable [34].
Despite the apparent support for the use of family guilds as
surrogates for species guilds, exceptions are obvious. The
difference in Gerês in the abundance of specialists between family
and species-level guilds is mainly due to the high abundance of
Dipoena melanogaster (C.L. Koch, 1837), the most commonly
sampled species [30]. Although this theridiid was assigned to the
space web builders’ guild according to its family, D. melanogaster is a
webless stenophagous ant specialist. At the global level, the bolas
spiders (genera Cladomelea, Mastophora and Ordgarius) are also
exceptions because these specialized hunters construct ‘‘bolas’’
that depend on aggressive chemical mimicry of a few moth species
rather than typical orb webs [52]. The triangular araneid spiders
Arkys have abandoned web building altogether and ambush prey
with large front legs like those of thomisids. Cybaeidae usually
build sheet-webs, but the water spider Argyroneta aquatica (Clerck,
1757) builds a silk retreat under water where it hunts without using
a web during the entire life cycle. Although many pisaurids are
active hunters, most spin large webs, while Dolomedes hunts on the
water surface, occasionally for small fish (in fact, the monophyly of
all Pisauridae is questionable). The same pattern repeats within
smaller lineages recognized as genera. Anapistula ataecina Cardoso &
Scharff, 2009 spin sheet-webs, not typical Anapistula orbs, probably
because of their subterranean habitat [53]. Even with many
exceptions, members of the same family do tend to present similar
ecological characteristics; hence taxonomical affiliation often is
associated with guild affiliation and the high correlation values
found by this study. The guilds suggested here can be applied with
care in many studies at various geographic scales.
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Proportions of guilds per site necessarily depend on the
sampling methods used. Most of our study datasets lacked data
for pitfall traps, even though the method often captures more
epigean fauna than any other method [30,31,33,37], and therefore
we could not properly assess ground hunters across datasets. The
precise proportions per guild (and family) found are therefore
specific to the methods available for comparison.
As predicted, guild composition was more stable than
taxonomic composition, suggesting turnover in families using
similar resources in different regions. Pholcids are relatively rare in
the temperate and Mediterranean faunas, but common through-
out the tropics and even dominant in the Tanzanian site.
Linyphiids are much more abundant in temperate regions than
elsewhere in the subtropics and tropics (except Tanzania, where
they are atypically diverse). Other families show this pattern of
high abundance in some regions but relative rarity in other regions
(e.g. Salticidae, Agelenidae, Theridiosomatidae, Philodromidae
and Oxyopidae).
Higher family richness in the tropics but similar functional
diversity suggest either greater functional role redundancy
compared to temperate forests or finer resource partitioning. In
a community with higher redundancy, the role of any one taxon
may be at least partly compensated by another taxon as the niches
of syntopic taxa present higher overlap and therefore it is easier for
the assemblage to keep its structure under disturbance and harder
for invading species to occupy ‘‘empty’’ niches. High ecological
redundancy may underlie the resilience of ecosystems to
disturbance and invasive species [40,54–56]. Assemblages with
particularly low functional diversity and very simple food chains,
such as sites in extreme altitudes or latitudes, small isolated oceanic
islands and caves, may be more susceptive to disruption [57].
Compared to other assemblages, these four examples may present:
1) low redundancy and therefore greater susceptibility to
disturbance; and 2) no suitable refuge from such disturbance.
Such assemblages are thus particularly vulnerable to habitat
destruction, invasive species and climate change. This diversity-
stability relation may explain why oceanic island assemblages, with
few species and sparse guilds, are especially prone to extinctions,
including for spiders [17]. The patterns found in this work may
also be partly explained by finer resource partitioning instead of or
besides redundancy. In the tropics many species can present
higher specialization in response to higher competition from
syntopic taxa. This results in narrower niches and higher number
of species or higher taxa per guild.
Functional diversity in Tanzania and Madagascar was less than
in other tropical sites and not statistically different from the
Portugal (Arrábida) site. These two tropical sites are however at
mid to high altitude. Comparing the three sites in temperate/
Mediterranean regions, at similar latitudes, the same tendency is
present, with the mid-altitude sites presenting a lower functional
diversity than the low-altitude site. If altitude correlates to some
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